New national and regional bryophyte records, 46

Abstract

Bryophyte biomass and diversity vary strongly with altitude in the tropics. Low abundance and low species numbers in lowland rain forests are most likely due to reduced diurnal activity times combined with high nocturnal respiration rates at high temperatures. This may exclude many montane species from the warm lowlands. However, an alternative hypothesis explains the observed pattern, namely a limited desiccation tolerance of montane species, precipitation being more concentrated but less frequent in most lowland forests compared to montane cloud forests. To test this hypothesis, we studied the desiccation tolerance of four montane and four lowland bryophyte species. The effects of prolonged drought were quantified with chlorophyll fluorescence (F_v/F_m) and the extent of electrolyte leakage. Both montane and lowland species survived dry periods of ≧80 days, which far exceeds the duration of dry periods in the wet lowland tropics. We can thus exclude intolerance to long dry spells as an explaination for the absence of the tested montane species in the lowlands. We should continue to focus on other mechanisms to explain the altitudinal gradient of bryophyte abundance and diversity in the tropics, in order to understand this pattern, as well as to predict future trends under climatic warming.     

Is hemiepiphytism an adaptation to high irradiance? Testing seedling responses to light levels and drought in hemiepiphytic and non‐hemiepiphytic Ficus

The epiphytic growth habit in many Ficus species during their juvenile stages has commonly been hypothesized to be an adaptation for avoiding deep shade in the forest understory, but this has never been tested experimentally. We examined growth and ecophysiology in seedlings of three hemiepiphytic (Hs) and three non‐hemiepiphytic (NHs) Ficus species grown under different irradiance levels. Both Hs and NHs exhibited characteristics of high light requiring species, such as high plasticity to growth irradiance and relatively high maximum photosynthetic assimilation rates. Diurnal measurements of leaf gas exchange showed that Hs have much shorter active photosynthetic periods than NHs; moreover, leaves of Hs have lower xylem hydraulic conductivity but stronger drought tolerance as indicated by much lower rates of leaf diebacks during the drought treatment. Seedlings of NHs had 3.3‐ and 13.3‐fold greater height and biomass than those of Hs species after growing in the nursery for 5 months, indicating a trade‐off between growth and drought tolerance due to the conflicting requirements for xylem conductivity and cavitation resistance. This study does not support the shade‐avoidance hypothesis; rather, it suggests that the canopy regeneration in Hs is an adaptation to avoid alternative terrestrial growth‐related risks imposed to tiny Ficus seedlings. The NHs with terrestrial regeneration reduce these risks by having an initial burst of growth to rapidly gain relatively large seedling sizes, while in Hs seedlings more conservative water use and greater drought tolerance for surviving the canopy environment are intrinsically associated with slow growth.     

Provenance variation in morphology and leaflet anatomy of Parkia biglobosa and its relation to drought tolerance

 Measurements of morphological and anatomical characteristics made on 4-month-old seedlings from five provenances of Parkia biglobosa (Jacq.) Benth., grown in a tropical greenhouse under controlled conditions, showed significant differences between the northern provenances, which represented the Sudan savanna zone, and the southern provenances which represented the forest zone of the species distribution. Seedlings of provenances from the savanna zone showed more xerophytic characteristics than those from the forest zone. They were smaller in height, with a greater number of smaller leaflets per pinnae, a lower specific leaflet mass, smaller palisade and guard cells and a lower ratio of palisade:non-palisade tissue. There were also significant differences in the same characteristics, except palisade cell length, between provenances from the easterly and westerly extremes of the species range. Stomatal conductance measured after 2 weeks of droughting also showed that provenances from the savanna zone had higher values, indicating greater physiological activity and reduced stress, compared with provenances from the forest zone. Multivariate discriminant analysis of the morphological and anatomical characteristics showed that 98% of the seedlings were classified correctly with respect to provenance. The squared distances between provenances were closely related to the geographical distances between locations. Yet, not all variation in leaf anatomy could be explained by the difference in climatic conditions from north to south. Also drought tolerance was not always related to morphological and anatomical features. It is possible that clinal variation within the species from east to west may be interacting with climatic differences from north to south. Received: 2 June 1998 / Accepted: 29 June 1998     

Germination and seedling frost tolerance differ between the native and invasive range in common ragweed

Germination characteristics and frost tolerance of seedlings are crucial parameters for establishment and invasion success of plants. The characterization of differences between populations in native and invasive ranges may improve our understanding of range expansion and adaptation. Here, we investigated germination characteristics of Ambrosia artemisiifolia L., a successful invader in Europe, under a temperature gradient between 5 and 25 °C. Besides rate and speed of germination we determined optimal, minimal and maximal temperature for germination of ten North American and 17 European populations that were sampled along major latitudinal and longitudinal gradients. We furthermore investigated the frost tolerance of seedlings. Germination rate was highest at 15 °C and germination speed was highest at 25 °C. Germination rate, germination speed, frost tolerance of seedlings, and the temperature niche width for germination were significantly higher and broader, respectively, for European populations. This was partly due to a higher seed mass of these populations. Germination traits lacked evidence for adaptation to climatic variables at the point of origin for both provenances. Instead, in the native range, seedling frost tolerance was positively correlated with the risk of frosts which supports the assumption of local adaptation. The increased frost tolerance of European populations may allow germination earlier in the year which may subsequently lead to higher biomass allocation—due to a longer growing period—and result in higher pollen and seed production. The increase in germination rates, germination speed and seedling frost tolerance might result in a higher fitness of the European populations which may facilitate further successful invasion and enhance the existing public health problems associated with this species.     

Geographic origin and past climatic experience influence the response to late spring frost in four common grass species in central Europe

Late spring frost events can affect vegetation. The response of grassland species, however, is generally unknown. We explore the late‐frost sensitivity of four common European grass species and investigate whether these species exhibit local adaptations to late frost on a continental scale and whether past climatic experience influences late frost sensitivity. Ecotypes of Arrhenatherum elatius, Alopecurus pratensis, Festuca pratensis, and Holcus lanatus from Spain, Italy, Bulgaria, Hungary, Sweden and Germany were exposed to late frost after drought and warming manipulations in the preceding growing season in a common garden experiment. Late frost reduced the productivity of the grasses on average by 20%. Ecotypes differed in their late‐frost sensitivity in three of the four species and local adaptations to late frost were identified. Previous exposure to drought and warming caused differences in late‐frost sensitivity in some cases. The impact of late frost events may increase in a warmer world due to an earlier onset of growing and no change in timing of late frost events. The history of climatic exposure can alter the performance of plants, possibly through epigenetic mechanisms. Based on the complex response pattern observed, a maximization of genetic diversity is proposed as a promising adaptation strategy against climate change.     

Functional traits predict drought performance and distribution of Mediterranean woody species

Water availability is one of the key environmental factors that affect plant establishment and distribution. In many regions water availability will decline with climate change, exposing small seedlings to a greater likelihood of drought. In this study, 17 leaves, stem, root, and whole-plant traits of ten woody Mediterranean species were measured under favourable growing conditions and seedling drought survival was evaluated during a simulated dry-down episode. The aims of this study were: i) to assess drought survival of different species, ii) to analyse which functional traits predict drought survival time, and iii) to explain species distribution in the field, based on species drought survival and drought strategies. Drought survival time varied ten-fold across species, from 19 to 192 days. Across species, drought survival was positively related to the rooting depth per leaf area, i.e., the ability to acquire water from deeper soil layers while reducing transpiring leaf area. Drought survival time was negatively related to species ability to grow quickly, as indicated by high relative growth and net assimilation rates. Drought survival also explained species distribution in the field. It was found that species were sorted along a continuum, ranging between two contrasting species functional extremes based on functional traits and drought performance. One extreme consisted of acquisitive fast-growing deciduous species, with thin, soft metabolically active leaves, with high resource use and vulnerability to drought. The opposite extreme consisted of conservative slow-growing evergreen species with sclerophyllous leaves, deep roots, a low transpiring area, and low water use, resulting in high drought survival and drought tolerance. The results show that these drought strategies shape species distribution in this Mediterranean area.     

Colonization with arbuscular mycorrhizal fungus affects growth,drought tolerance and expression of stress-responsive genes in <Emphasis Type="Italic">Poncirus trifoliata</Emphasis>

The objective of this study was to investigate the effects of arbuscular mycorrhizal fungus (AMF) inoculation on growth and drought tolerance of Poncirus trifoliata seedlings. The seedlings were inoculated with or without Glomus mosseae before exposure to a short-term (3 days) water depletion, and relevant physiological and biochemical parameters (plant height, chlorophyll content, relative water content, activity of antioxidant enzymes) and expression patterns of several stress-responsive genes were examined. Inoculation with G. mosseae led to growth promotion of the seedlings, as revealed by larger plant height and higher relative water and chlorophyll contents. When subjected to drought treatment, the AMF-inoculated (AM) plants showed better tolerance than the nonmycorrhizal (NAM) plants. Under drought, the AM plants exhibited higher level of proline and activity of two antioxidant enzymes, superoxide dismutase (SOD) and peroxidase (POD). In addition, mRNA abundance of four genes involved in reactive oxygen species homeostasis and oxidative stress battling was higher in the AM plants when compared with the NAM plants. These results indicate that AMF inoculation stimulated growth and enhanced drought tolerance of the seedlings, which may be due to activation of an arsenal of physiological, biochemical and molecular alterations.     

Contrasting climate influences on Nothofagus pumilio establishment along elevational gradients

Predicted warmer temperatures and more frequent extreme climatic events in the southern Andes may affect the dynamics of the Patagonian forests. These environmental changes may differentially alter the probability of Nothofagus pumilio establishment across its altitudinal range of distribution. We monitored fruit fall, seedling emergence and survival at altitudinal distribution range of N. pumilio forests in Santa Cruz (49°22′ S—72°56′ W), Argentina. Fruit fall, seedling emergence and survival were tested in relation to drought, based on the SPEI (Standardized Precipitation Evapotranspiration Index), interacting with elevation. Fruit fall was consistently higher at low elevation and the abundance of fruits was not affected by drought intensity. Density of new-born seedlings was?~?9 to 24 times lower at low- than at high-elevation plots in our first observations (2014 and 2016), characterized by warm-dry climatic conditions in spring-early summers. In contrast, seedling abundance was?~?1.5 times larger at low elevations during relatively cold-wet growing seasons. Survival probability was explained by the interaction between SPEI and elevation. At low elevation and in dry periods, survival probability was lower (CI 54–72%) than in wet periods (CI 68–84%) but at high elevation similar survival was registered even with positive or negative SPEI values. Our results show interacting effects of elevation and drought on tree establishment at the elevation limits, with positive and negative drought effects at high and low elevations, respectively. Predicted increase of extreme drought events during the XXI century could be detrimental for N. pumilio establishment at dry, low-elevation forests.

     

Can environmental tolerances explain convergent patterns of distribution in endemic spring snails from opposite sides of the Australian arid zone?

Patterns of distribution are influenced by species environmental requirements and limits, but experimental tests are needed to discern whether correlates of abundance directly affect survival and success. Springs in Australia’s arid interior support a wide diversity of gastropods only found in springs, and these species show dichotomous patterns of distribution. “Amphibious” species are broadly distributed across many springs and microhabitats, and “aquatic” species confined to the deepest pool areas within large springs. This pattern appears to be driven by the interaction between different environmental conditions in different microhabitats and the environmental tolerances of each endemic snail species. Factorial experiments were used to test whether conditions in the environmentally extreme and variable tail area of springs (considering pH, conductivity, temperature and desiccation potential, alone and in synergistic scenarios) elicited lethal or sub-lethal responses in spring snails endemic to springs on opposite sides of the Australian arid zone. All species restricted to spring pools were able to endure 24 h exposed to the average tail conditions, alone and in combination, but most suffered mortalities when subjected to extremes, and mortalities occurred sooner in the most restricted species when elevated pH and conductivity were experienced in combination. Responses of species from different locations are similar, but pattern of distribution in the field were not correlated with tolerance of environmental extremes—with the “amphibious” species from the sub-tropics being far more sensitive than its arid counterpart. These findings suggest that environmental variance within springs can influence patterns of distribution and abundance, particularly when extremes are experienced simultaneously over sustained time periods. But despite similarities in responses across species from these two spring complexes, no simple generalisations linking distribution and tolerance were discernible.     

Tempo and mode of the multiple origins of salinity tolerance in a water beetle lineage

Salinity is one of the most important drivers of the distribution, abundance and diversity of organisms. Previous studies on the evolution of saline tolerance have been mainly centred on marine and terrestrial organisms, while lineages inhabiting inland waters remain largely unexplored. This is despite the fact that these systems include a much broader range of salinities, going from freshwater to more than six times the salinity of the sea (i.e. >200 g/L). Here, we study the pattern and timing of the evolution of the tolerance to salinity in an inland aquatic lineage of water beetles (Enochrus species of the subgenus Lumetus, family Hydrophilidae), with the general aim of understanding the mechanisms by which it was achieved. Using a time‐calibrated phylogeny built from five mitochondrial and two nuclear genes and information about the salinity tolerance and geographical distribution of the species, we found that salinity tolerance appeared multiple times associated with periods of global aridification. We found evidence of some accelerated transitions from freshwater directly to high salinities, as reconstructed with extant lineages. This, together with the strong positive correlation found between salinity tolerance and aridity of the habitats in which species are found, suggests that tolerance to salinity may be based on a co‐opted mechanism developed originally for drought resistance.     

Drought effects on seedling survival in a tropical moist forest

The amount and seasonality of rainfall varies strongly in the tropics, and plant species abundance, distribution and diversity are correlated with rainfall. Drought periods leading to plant stress occur not only in dry forests, but also in moist and even wet forests. We quantified experimentally the effect of drought on survival of first year seedlings of 28 co-occurring tropical woody plant species in the understory of a tropical moist forest. The seedlings were transplanted to plots and subjected to a drought and an irrigation treatment for 22 weeks during the dry season. Drought effects on mortality and wilting behavior varied greatly among species, so that relative survival in the dry treatment ranged from 0% to about 100% of that in the irrigated treatment. Drought stress was the main factor in mortality, causing about 90% (median) of the total mortality observed in the dry treatment. In almost half of the species, the difference in survival between treatments was not significant even after 22 weeks, implying that many of the species are well adapted to drought in this forest. Relative drought survival was significantly higher in species associated with dry habitats than in those associated with wet habitats, and in species with higher abundance on the dry side of the Isthmus of Panama, than in those more abundant on the wet side. These data show that differential species survival in response to drought, combined with variation in soil moisture availability, may be important for species distribution at the local and regional scale in many tropical forests.     

Could phenotypic plasticity limit an invasive species? Incomplete reversibility of mid-winter deacclimation in emerald ash borer

The emerald ash borer (Agrilus planipennis, Coleoptera: Buprestidae) is a wood-boring invasive pest devastating North American ash (Fraxinus spp.). A. planipennis overwinters primarily as a freeze-avoiding prepupa within the outer xylem or inner bark of the host tree. The range of this species is expanding outward from its presumed introduction point in southwestern Michigan. We hypothesized that loss of cold tolerance in response to mid-winter warm spells could limit survival and northern distribution of A. planipennis. We determined whether winter-acclimatised A. planipennis prepupae reduced their cold tolerance in response to mid-winter warm periods, and whether this plasticity was reversible with subsequent cold exposure. Prepupae subjected to mid-winter warm spells of 10 and 15°C had increased supercooling points (SCPs) and thus reduced cold tolerance. This increase in SCP was accompanied by a rapid loss of haemolymph cryoprotectants and the loss of cold tolerance was not reversed when the prepupae were returned to −10°C. Exposure to temperatures fluctuating from 0 to 4°C did not reduce cold hardiness. Only extreme warming events for several days followed by extreme cold snaps may have lethal effects on overwintering A. planipennis populations. Thus, distribution in North America is likely to be limited by the presence of host trees rather than climatic factors, but we conclude that range extensions of invasive species could be halted if local climatic extremes induce unidirectional plastic responses.     

Interspecific variation in functional traits of oak seedlings (<Emphasis Type="Italic">Quercus ilex,Quercus trojana,Quercus virgiliana</Emphasis>) grown under artificial drought and fire conditions

To face summer drought and wildfire in Mediterranean-type ecosystems, plants adopt different strategies that involve considerable rearrangements of biomass allocation and physiological activity. This paper analyses morphological and physiological traits in seedlings of three oak species (Quercus ilex, Quercus trojana and Quercus virgiliana) co-occurring under natural conditions. The aim of this study was to evaluate species-specific characteristics and the response of these oak seedlings to drought stress and fire treatment. Seedlings were kept in a growth chamber that mimicked natural environmental conditions. All three species showed a good degree of tolerance to drought and fire treatments. Differences in specific biomass allocation patterns and physiological traits resulted in phenotypic differences between species. In Q. ilex, drought tolerance depended upon adjustment of the allocation pattern. Q. trojana seedlings undergoing mild to severe drought presented a higher photosystem II (PSII) efficiency than control seedlings. Moreover, Q. trojana showed a very large root system, which corresponded to higher soil area exploitation, and bigger leaf midrib vascular bundles than the other two species. Morphological and physiological performances indicated Q. trojana as the most tolerant to drought and fire. These characteristics contribute to a high recruitment potential of Q. trojana seedlings, which might be the reason for the dominance of this species under natural conditions. Drought increase as a result of climate change is expected to favour Q. trojana, leading to an increase in its spatial distribution.     

Abiotic constraints on the establishment of Quercus seedlings in grassland

High evaporative demand and periodic drought characterize the growing season in midwestern grasslands relative to deciduous forests of the eastern US, and predicted climatic changes suggest that these climatic extremes may be exacerbated. Despite this less than optimal environment for tree seedling establishment, deciduous trees have expanded into adjacent tallgrass prairie within the last century leading to a dramatic land cover change. In order to determine the role of light and temperature on seedling establishment, we assessed carbon and water relations and aboveground growth of first‐year Quercus macrocarpa seedlings exposed to one of three conditions: (1) intact tallgrass prairie communities (control), (2) aboveground herbaceous biomass removed (grass removal), and (3) shade plus biomass removal to reduce light (PFD) to levels typical of the grassland‐forest ecotone (shade). In the 2000 growing season, precipitation was 35% below the long‐term average, which had a significant negative effect on oak seedling carbon gain at midseason (photosynthesis declined to 10% of maximum rates). However, net photosynthesis and stomatal conductance in the shade treatment was ca. 2.5 and 1.5 times greater, respectively, than in control treatment seedlings during this drought. During this period, leaf and air temperatures in control seedlings were similar whereas leaf temperatures in the shade treatment remained below air temperature. A late‐season recovery period, coincident with decreased air temperatures, resulted in increased net photosynthesis for all seedlings. Increased photosynthetic rates and water relations in shaded seedlings compared to seedlings in full sun suggest that, at least in dry years, high light and temperature may negatively impact oak seedling performance. However, high survival rates for all seedlings indicate that Q. macrocarpa seedlings are capable of tolerating both present‐day and future climatic extremes. Unless historic fire regimes are restored, forest expansion and land cover change are likely to continue.     

Seedling Traits Determine Drought Tolerance of Tropical Tree Species

Water availability is the most important factor determining tree species distribution in the tropics, but the underlying mechanisms are still not clear. In this study, we compared functional traits of 38 tropical tree species from dry and moist forest, and quantified their ability to survive drought in a dry‐down experiment in which wilting and survival were monitored. We evaluated how seedling traits affect drought survival, and how drought survival determines species distribution along the rainfall gradient. Dry forest species tended to have compound leaves, high stem dry matter content (stem dry mass/fresh mass), and low leaf area ratio, suggesting that reduction of transpiration and avoidance of xylem cavitation are important for their success. Three functional groups were identified based on the seedling traits: (1) drought avoiders with a deciduous leaf habitat and taproots; (2) drought resisters with tough tissues (i.e., a high dry matter content); and (3) light‐demanding moist forest species with a large belowground foraging capacity. Dry forest species had a longer drought survival time (62 d) than moist forest species (25 d). Deciduousness explained 69 percent of interspecific variation in drought survival. Among evergreen species, stem density explained 20 percent of the drought survival. Drought survival was not related to species distribution along the rainfall gradient, because it was mainly determined by deciduousness, and species with deciduous seedlings are found in both dry and moist forests. Among evergreen species, drought survival explained 28 percent of the variation in species position along the rainfall gradient. This suggests that, apart from drought tolerance, other factors such as history, dispersal limitation, shade tolerance, and fire shape species distribution patterns along the rainfall gradient.     

Climate and plant distribution at global and local scales

This paper investigates, with predictive models, the utility of ecophysiological responses to climate as predictors of plant distribution. At the global scale responses to extreme minimum temperatures and to the hydrological budget effectively predict the distribution limits of the major vegetation types of the World. A minimum temperature of -15°C, for example, appears critical in controlling the poleward spread of vegetation that is dominated by evergreen broadleaved species; however, the presence or absence of more frost resistant species, such as those that are deciduous broadleaved, is not obviously explained in terms of extremes of climate. In such cases, predicting the competitive relationships between species is necessary and dependent on the climatic sensitivity of population dynamics.We are grateful to Hans de Kroon for discussions on the application of matrices to ecology.     

Biogeography of Banksia in southwestern Australia

Comprehensive new data bases on the taxonomy, distribution, ecology and climate of Banksia (Proteaceae) were used to study the biogeography of the sixty endemic species in southwestern Australia. They occur at the wettest coastal tip, with up to 1.5 m rainfall p.a., 1500 km along the north coast, 1000 km along the south coast and up to 450 km inland to the desert margin, with as low as 0.2 m rainfall p.a. The largest areas (sum of 30´ lat.×30´ long. grid cells) containing banksias were under much more arid climates than those with highest species density. Species density was usually lowest at the climatic extremes. Two nodes of species richness and ten floristic groups, each with a characteristic climate, landscape, vegetation and soil types and suite of indicator and endemic species, were recognized. A case is made for resurrecting the Lesueur (northern node) and Stirling (west portion of the southern node) phytogeographic districts, together with separating the West (east portion of the southern node) from the East Eyre district. These species-rich areas coincide with nutrient impoverished sands and laterite associated with dissected uplands having low, highly sclerophyllous vegetation. The distribution of strongly serotinous species and those with persistent florets is consistent with greater reliance on fire for creating suitable recruitment conditions with increasing aridity. The distribution of needle-leaved species is only partly explained by their superior drought tolerance. A preliminary transplant trial with two Banksia species showed that their distribution was only partly restricted by their current climatic profile.     

Water availability,fundamental niches and realized niches: A case study from the Cape flora

The ability of plants to survive drought or waterlogging constitutes an important niche parameter, which might be particularly significant in explaining species coexistence in the species‐rich and seasonally dry Cape Floristic Region of South Africa. However, the degree of physiological adaptation and specialization to these eco‐hydrological parameters (the fundamental niche) cannot be readily inferred from correlative studies based on species distributions and spatial variation in environmental parameters (the realized niche). We used an ex situ greenhouse experiment to compare the fundamental hydrological niches (different mean annual precipitation, rainfall seasonality and soil drainage) of six eco‐hydrologically divergent African Restionaceae species. Juvenile plants were subjected to six different watering treatments, ranging from no watering to waterlogging, to determine drought and waterlogging susceptibility and optimal growth conditions. We used the rate of biomass accumulation and survival rate as response measures. We found that species from dry and mesic (but well‐drained) habitats had optimal or near‐optimal growth at benign conditions (under which most restio species grow well). All species performed worse when droughted and died when not watered. Species from dry habitats tended to perform better (assessed in growth) than species from wet habitats under droughting. Species from wet habitats performed best when waterlogged, whereas species from dry habitats performed very poorly when waterlogged – thus showing that realized and fundamental niches covaried at the wet end of the hydrological gradient. We conclude that eco‐hydrological parameters are part of the fundamental niche, and fundamental and realized species niches are approximately correlated along them. The distribution of wet habitat species appears not to reflect their drought tolerance, suggesting that it may not be predicted by bioclimatic variables, but rather by soil drainage characteristics.